PROCEEDINGS

OF THE

FORTEENTH SYMPOSIUM

ON THE

NATURAL HISTORY OF THE BAHAMAS

Edited by Craig Tepper and Ronald Shaklee

Conference Organizer Thomas Rothfus

Gerace Research Centre San Salvador Bahamas 2011

Cover photograph – “Iggie the Rock Iguana” courtesy of Ric Schumacher

Copyright Gerace Research Centre

All Rights Reserved

No part of this publication may be reproduced or transmitted in any form or by any means electronic or mechanical, including photocopy, recording or information storage or retrieval system without permission in written form.

Printed at the Gerace Research Centre

ISBN 0-935909-95-8

The 14th Symposium on the Natural History of the Bahamas

GENETIC DIVERSITY OF THE BLUE LAND , GUANHUMI LATREILLE (CRUSTACEA: ), ON ANDROS ISLAND: IMPLICATIONS FOR SUSTAINABLE MANAGEMENT

Tami L. LaPilusa and Jeffrey S. Heilveil Biology Department State University of New York, College at Oneonta 108 Ravine Parkway Oneonta, NY 13820

ABSTRACT INTRODUCTION

Androsians provide for their families by Human populations have traditionally sub- harvesting marine and terrestrial organisms on sisted on organisms harvested within the local en- and around the island. These , including vironment (Boone, 2002). Edible species of great , the blue land crab, provide supply are often the target of regular, small-scale not only food, but also a substantial supplement to artisanal fishery harvesting (Trosper, 2002). Even- the annual economy of Andros Island (Rolle, tual commercialization of artisanal fisheries has 2007; ANCAT, 2010). As habitat fragmentation led to collapses in fishery stocks, as occurred with and harvesting pressure increase, many species the anchovy (Tarazona, 1983) and is occurring have declined or been extirpated. It is becoming with the Nassau grouper (Sadovy and Eklund increasingly important to understand the genetic 1999; ANCAT, 2011; Cornish and Eklund, 2003). diversity of these cottage industry species. Andros Seemingly limitless resources provide a false Island is known to have the largest sense of resource and economic security leading metapopulation of blue land in The Baha- to a tragedy (sensu Hardin, 1968) when human mas (ANCAT, 2010), providing generations of use is not balanced with population sustainability Androsians with income derived from the bounti- of commodity species. ful harvest during the spawning migrations. Lo- The islands of The Bahamas have been in- cals have reported having to travel further from habited by humans since around 600 A.D. (Craton their settlements to collect adequate numbers of and Saunders, 1992). Fishery resources have been, crabs to send to market, due to increased harvest- and continue to be, an important source of food ing pressure. This project analyzed genetic varia- and revenue for the island nation. The spiny lob- tion at the nuclear arginine kinase gene and mi- ster and Nassau grouper fisheries that grew out of crosatellite markers to identify the discrete popu- early artisanal small-scale harvesting now make lations of C. guanhumi and quantify the extent of up more than 75% of the commercial fishery re- its genetic diversity on Andros Island. Preliminary sources with an estimated annual economic con- analysis of arginine kinase sequences revealed tribution of $105M USD (Rankine et al., 2004). limited genetic diversity between populations in Most of the commercial fisheries in The Bahamas the less-settled portions of Andros. Mapping the are based out of the main population centers on extent of the populations and amount of genetic Grand Bahama and New Providence. Historically, diversity in each will allow scientific data to be the people living on the sparsely populated islands used to improve management strategies to con- continued to harvest in much smaller quantities, serve both C. guanhumi and its economic value to putting less pressure on fishery stocks. Increases Androsians. in tourism on these less well-known islands have created a market for increased harvesting of the artisanal fishery species (Danylchuk, 2005). Cardisoma guanhumi Latrielle, the blue land crab,

41 The 14th Symposium on the Natural History of the Bahamas is emerging as a profitable fishery resource as the Formal studies are not readily available smaller island communities struggle to remain regarding C. guanhumi on Andros Island and even economically viable. The large numbers of blue the number of populations on the Island is un- land crabs on Andros Island are becoming well- known. This study used nuclear and microsatellite known and increasing numbers of land crabs are markers to determine the population structure of being collected for use in tourist-based heritage C. guanhumi on Andros Island, The Bahamas, festivals across The Bahamas (Miller, 2005; DCL, with the goal of long-term sustainability and sci- 2005). ence-based conservation to preserve both the crab Andros Island is believed to have the larg- and its economic value to Androsians. est population of Cardisoma guanhumi in The Bahamas and Androsians are known throughout the islands for their crab collecting expertise (ANCAT, 2010; Rolle, 2007). Individuals have been known to collect over 100 crabs in an even- ing; with groups of crabbers quickly collecting dozens in a short sweep of the migratory path (Hostetler et al., 1991; Wedes, 2004; D. Scott, pers. comm.). The blue land crab provides a good source of income, fetching up to $5 per crab at market, depending on the season and the fat con- tent of the crab. This high price allows seasoned collectors to add an estimated $20,000 to their an- nual income in just 2-3 months (BIS, 2002). The Figure 1. Cardisoma guanhumi, ovigerous female artisanal crabbing industry has allowed homes to be built and provided college tuition for many MATERIALS AND METHODS Androsian students (ANCAT, 2010). The maintenance of genetic diversity is Study Organism emerging as a new paradigm for managing com- modity organisms to promote economic prosperity Cardisoma guanhumi Latreille (Crustacea: in developing nations (e.g. Dawson et al., 2009). Decapoda; Figure 1) is a land crab found through- Conservation genetics uses genetic information to out the Atlantic coastal sub-tropic and tropic estu- assess populations under threat of habitat loss, arine areas from Florida to Brazil (Burggren and exploitation and/or environmental change McMahon, 1988). Juvenile crabs develop in man- (Frankham, 2002). The cornerstone of this type of grove estuaries and, with the onset of sexual ma- management rests on a good understanding of the turity around four years of age, adults create bur- underlying genetic structure of the species in rows in the hardwood coppices (Hostetler et al., question. DeOliveria-Neto et al. (2008) found a 1991). Spawning migrations typically start at the high degree of genetic diversity throughout the beginning of the rainy season (Witherington, blue land crab populations along the coast of Bra- 2009) which varies throughout the range based on zil. latitude (Wedes, 2004). Females will carry a Years of overharvesting (Amaral and clutch of 300,000 - 700,000 fertilized eggs for Jablonski, 2005) and mangrove removal (Magris about 2 weeks while development occurs and Barreto, 2010) have put additional pressure (Hostetler et al., 1991) until the hatching larvae on those populations. The Brazilian Ministry of can be released into the sea. Major and minor mi- Environment includes C. guanhumi as an over- gratory waves occur during the full and new harvested species in spite of regulations limiting moons (Gifford, 1962; Hill, 2001), during which harvest to only males with carapace length greater the females are known to travel up to 8 Km to than 4.5 cm (Alves et al., 2005). shed the hatching larvae into the sea (Burggren

42 The 14th Symposium on the Natural History of the Bahamas and McMahon, 1988). Adult females will return performed by hand. Traditional analyses to their burrows and await the next annual migra- (AMOVA, FST, Pairwise differentiation) were cal- tory spawn (Burggren and McMahon, 1988). Fe- culated with Arlequin 3.5 (Ver 3.5, Excoffier et males can reproduce 11 times, on average, during al., 2005). a life-span of approximately 15 years (Witherington, 2009).

Sample collection and DNA Extraction

Geographic clines and indigenous knowledge were utilized to select thirteen poten- tial sites on Andros Island, The Bahamas (Figure 2). Live crabs (n = 3-20) were collected by hand and a single pereopod was removed and preserved in 95% ethanol. Crabs were subsequently released as close to the point of collection as possible. Muscle was removed from each of the preserved pereopods and genomic DNA was extracted (DNeasy Blood and Tissue Kit, Qiagen, Valencia, CA). Samples were stored at -80o C and vouch- ered with the State University of New York, Col- lege at Oneonta’s Biological Field Station.

Arginine Kinase gene amplification, sequencing, and analysis

A 321-base region of arginine kinase was amplified using the AKF3 and AKR3 primers of Mahon and Neigel (2008). Amplification consist- Figure 2. Andros Island Collection Site Map ed of initial denaturation at 95o C for 2 min, fol- (modified from Nickrent et. al., 1988). Number of lowed by 35 cycles of denaturation at 95o C for 45 individuals sampled for each site indicated in pa- sec, annealing at 75o, 70o, or 67o C (depending on rentheses after site identification number. individual) for 1 min, extension of 72o C for 1 min, and a final extension at 72o C for 5 min. Microsatellite PCR and Fragment Analysis Each 25 µL PCR was prepared according to the instructions of the manufacturer (5 Prime, Microsatellite loci TJH-06 of Zhang et al. Gaithersburg, MD); however, only 1 unit of (2009) and pm-109 of Fratini et al. (2006) were HotMaster™ Taq DNA Polymerase was used. genotyped to assess genetic differentiation. Each After purification (QIAquick PCR Purification 25 µL PCR contained the following concentra- Kit, Qiagen, Valencia, CA), successful amplicons tions: 1 unit of HotMaster™ Taq DNA Polymer- ase, 1X HotMaster™ Taq Buffer with 25 mM were sequenced on a CEQ8000 (Beckman Coul- 2+ ter, Fullerton, CA). Mg , 0.5 µM dNTPs, 0.5µM unlabeled reverse Sequences were aligned using ClustalW primer, 0.5 µM dye-labeled forward primer, and 30 ng genomic DNA. The thermal cycler profile Multiple Sequence Alignment and the open read- o ing frame determined utilizing SIXFRAME via consisted of an initial denaturation of 95 C for 3 Bioworkbench (http://workbench.sdsc.edu/). Tests for Hardy-Weinberg Equilibrium (HWE) were

43 The 14th Symposium on the Natural History of the Bahamas

Table 1. Pairwise FST with corresponding p-value. FST values above the diagonal, associated p-values below the diagonal. Site numbers as per Figure 2. min; followed by 35 cycles of denaturation of lack of genetic differentiation. That the whole 95o C for 45 sec, annealing at 57o C for 1 min, island is in HWE for both expressed and unex- and extension of 72o C for 1 min; and a final pressed markers further suggests a single undif- extension at 75o C for 15 min. Fragments were ferentiated population. Preliminary analyses of analyzed on a CEQ8000 (Beckman Coulter, two microsatellite loci also supported lack of Fullerton, CA) utilizing 0.5 µL of 600 size- diversity, as both loci were monomorphic across standard, as per the instructions of the manufac- Andros Island. turer. Anecdotal evidence suggests that the blue land crab population has been steadily de- RESULTS clining over the last several decades. Daytime sightings along the spawning path have become Initial sequencing allowed analysis of a rare in the last decade (G and A. Colebrooke, 321-base fragment of arginine kinase for 72 in- pers. comm.). dividuals across Andros. Allelic richness for the The cyclical nature of blue land crab mi- island was three, with two variable sites. Two grations has historically been a time of intense rare genotypes were found in the southern por- collecting of C. guanhumi. The blue land crabs tions of North and South Andros Island (Sites 1 are collected on their way to the sea and upon and 9, respectively in Figure 2). Fragment anal- return to their burrows, with no regard to sex or ysis of 70 individuals for TJH-06 and 8 individ- reproductive status. Mangrove estuary habitat uals for pm-109 were monomorphic for a single has been eliminated to make way for exclusive allele at each locus across the entire island. All resorts and luxury home sites (pers. obs.). In populations were in HWE. The Analysis of Mo- 1998, the “All-Andros Crab Fest” was estab- lecular Variance (AMOVA) indicated that al- lished to celebrate the crabbing legacy of An- most all variation occurred within populations. dros Island and the “Opening of the Andros Pairwise FST values ranged from -0.187 to 0.186 Bank” (i.e. withdrawing the crabs from Andros but were not significantly different from zero as one would money from a savings account) (P-value = 0.09 - 0.99, Table 1). with 4 days of festivities high-lighted by a plethora of crab dishes and an influx of tourism DISCUSSION from throughout The Bahamas (BIS, 2002). People travel from other islands to collect crabs Preliminary analysis suggested C. guan- (pers. comm. G. Douglas) and anecdotal evi- humi on Andros Island comprise one population dence indicates there are no crabs to cook by the with limited genetic diversity. Both AMOVA last day of the festival, suggesting that the addi- and tests of pairwise differentiation support the

44 The 14th Symposium on the Natural History of the Bahamas

tive effect of changes in both habitat and collec- Local stakeholders will need to be con- tion has been disastrous. sulted in future management plans for the land Land Crab National Park was estab- crab throughout the range and more importantly, lished in 2002 on Andros as a means to preserve Andros. The “All-Andros Crabfest” provides a C. guanhumi habitat and migratory pathways prime opportunity to serve as a checkpoint dur- (BNT 2009), but its borders are not enforced, ing the harvest season to survey those involved since it was never the intention of the Bahamas in the industry to collect data on harvest num- National Trust (BNT) to limit land crab harvest bers and the dispersion of collection sites to in Land Crab National Park (S. Buckner, pers. manage the seasonality of the harvests. Sustain- comm.). able harvest education can also become a high- There is currently no season, limit, or light of the festival with the optimal placement size and/or sex restrictions on harvesting C. of the land crab education/conservation booths. guanhumi on Andros despite similar measures During the “All-Andros Crabfest 2011”, all ed- in other countries. Florida Fish and Wildlife cur- ucational displays were located in the far reach- rently regulates not only the allowable harvest es of Queen’s Park (pers. obs.) beyond where but also an established season for allowable col- most visitors would go. While it is important to lecting (Hostetler et al., 1991). Puerto Rico pro- highlight the habitat being protected, education hibits collection during the local spawning sea- of the local and tourist population is necessary son (Hostetler et al., 1991), though enforcement for successful management strategies. is ineffective (Govender, 2008) and harvesting Our study indicated a lack of genetic di- more intense and destructive outside of the versity in the C. guanhumi population on An- management areas (Rodríguez-Fourquet and dros. Further work with polymorphic markers Sabat, 2009). The Brazilian Ministry of the En- and on different islands in The Bahamas will vironment restricted collection of C. guanhumi assist in the establishment of Evolutionary Sig- with carapace widths less than 8 cm and estab- nificant Units (Crandall et al., 2000) to aid fu- lished an off season during the annual spawn ture conservation of the species through the (IBAMA, 2011, Ministério do Meio Ambiente, BNT. Any management strategy developed for 2011), although artisnal fishers continue to har- C. guanhumi must include sustainable harvest- vest without regard to the regulations put into ing with input from the local collectors. The place by the Ministry of the Environment (Alves combination of habitat preservation, resource et al., 2005). monitoring and conservation genetics utilized to Management strategies on Andros in- develop regulations may aid long-term sustaina- clude educational resources distributed through- bility of the blue land crab on Andros. out the local communities (Witherington, 2009) and through the BNT. Formal regulations have ACKNOWLEDGMENTS not been established through the BNT, though encouragement exists for collection of females We would like to thank the State Uni- only after eggs have been released into the versity of New York, College at Oneonta SUNY ocean (Witherington, 2009). Androsians indi- College Senate Committee on Research and The cate that females inadvertently caught are im- College at Oneonta Foundation for partial fund- mediately released due to lack of quality for ing of this research through two Student- consumption (A. Colebrooke, P. Riley, E. Wal- Initiated Research Grants awarded to T. lace, pers. comm.). Protection of habitat through LaPilusa. The College at Oneonta Alumni As- the Land Crab National Park continues; alt- sociation awarded an Alumni Scholarship to T. hough with no prohibition of harvest in the LaPilusa to further support this research. Addi- management areas, only habitat is being pro- tional financial support was provided by M. tected (BNT, 2009). LaPilusa. Logistical support was provided by

45

The 14th Symposium on the Natural History of the Bahamas

Dr. W. Ben Bohl and International Field Studies able at: www.bahamas.gov.bs. Online (IFS), Inc., Columbus, Ohio; and the IFS Forfar press release 6/14/02 [retrieved 3/21/10] Field Station, Blanket Sound, Andros Island, Bahamas provided accommodations and facili- BNT, 2009. Central Andros National Parks. tated travel on Andros Island. We would also Bahamas National Trust [online] Avail- like to thank Annie Colebrooke, Rivean Riley, able at: http://www.bnt.bs/parks_andros. Edricco Scott, Jazmin and Justin LaPilusa, and php?subid=17&catid=6 the interns at Forfar Field Station and IFS Blue [retreived 11/04/11] Water Sailing Program for assistance in the field. Collections were made under permits to JS Boone, J.L. 2002. Subsistence strategies and Heilveil and TL LaPilusa from The Bahamas early human population history. World Environment, Science, and Technology Com- Archaeology 34:6–25. mission, Ministry of the Environment and The Bahamas Department of Agriculture, Andros Burggren, W.W., and B.R. McMahon. 1988. Island, certificate # 01-1399-09 and United Biology of the Land Crabs. Cambridge. States Fish & Wildlife Service permit # Cambridge University Press, 479 pp. 2009MI781008. Cornish, A., and A.M. Eklund. 2003. REFERENCES Epinephelus striatus. in: IUCN 2010: IUCN Red List of Threatened Species. Alves, R.R.N., A.K. Nishida, and M.I.M. Her- Available at: www.iucnredlist.org/apps/ nández. 2005. Environmental perception redlist/details /7862/0 [retrieved 1/11/11] of gatherers of the crab 'caranguejo-uçá' ( cordatus, Decapoda, Brachyura) Crandall, K.A., R.P. Olaf, G. Bininda-Emonds, affecting their collection attitudes. Jour- M. Mace, and R. K. Wayne. 2000. Con- nal of Ethnobiology and Ethnomedicine sidering evolutionary processes in con- 1:10. servation biology. Trends in Ecology and Evolution 15: 290-295. Amaral, A.C.Z., and S. Jablonski. 2005. Con- servation of Marine and Coastal Biodiversity in Craton, M., and G. Saunders. 1992. Islanders in Brazil. Conservation Biology 19:625–631. the Stream: A History of the Bahamian People. Volume One: From Aboriginal ANCAT, 2010. Andros Crabs. Andros Conserv- Times to the End of Slavery. Athens: ancy and Trust [online] Available at: The University of Georgia Press, 455 pp. www.ancat.net/index.php?option=com_ content&view=article&id=61& Danylchuk, A.J. 2005. Fisheries management in Itemid=75 [retrieved 11/04/11] South Eluthera, Bahamas: Can a Marine reserve save the ‘Holy Trinity’?. 56th ANCAT, 2011. Spring 2011 Newsletter. Andros Gulf and Caribbean Fisheries Institute Conservancy and Trust [online] Availa- 56:169-177. ble at:http://www.ancat.net/ newsletters/ANCAT-Newslette- Dawson, I.K., A. Lengkeek, and J. C. Weber. AprilJune-2011.pdf [retrieved 11/05/11] 2009. Managing genetic variation in tropical trees: linking knowledge with BIS, 2002. Andros Crab fest excites patrons. action in agroforestry ecosystems for Bahamas Information Service. The improved conservation and enhanced Commonwealth of the Bahamas. Avail- livelihoods. Biodiversity Conservation

46

The 14th Symposium on the Natural History of the Bahamas

18:969-986. Govender, Y., A.M. Sabat, and E. Cuevas. 2008. Effects of land-use/ land-cover changes DCL, 2005. Grand Bahama Island Events: Mini on land crab, Cardisoma guanhumi, "Crab Fest". Discovery Cruise Line. abundance in Puerto Rico. Journal of [online]. Available at: www.discovery Tropical Ecology 24:417-423. cruiseline.com/ bahamas/events [retrieved 11/05/11] Hardin, G. 1968. The Tragedy of the Commons. Science 162:1243–1248. De Oliveria-Neto, J.F., M.R. Pie., M.A. Chammas, A. Ostrensky, and W.A. Hill, K. 2001. Cardisoma guanhumi. Smithson- Boeger. 2008. Phylogeography of the ian Marine Station at Fort Pierce blue land crab, Cardisoma guanhumi [online]. Available at: (Decapoda:) along the Bra- http://www.sms.si.edu/IRLFieldGuide/ zilian coast. Journal of the Marine Bio- Cardis_guanhu.htm [retrieved 1/26/09] logical Association of the United King- dom 88:1417-1423. Hostetler, M.E., F.J. Mazzotti, and A.K. Tayler. 1991. Blue Land Crab (Cardisoma Excoffier, L., L. Guillaume, and S. Schneider. guanhumi). WEC 30. Department of 2005. Arlequin (version 3.0): An inte- Wildlife Ecology and Conservation. grated software package for population Florida Cooperative Extension Service, genetics data analysis. Evolutionary Bio- Institute of Food and Agricultural Sci- informatics Online 1:47-50. ences, University of Florida. Available at: www. edis.ifas.ufl.edu/uw013 Frankham, R. 2002. Conservation Genetics. [retreived 1/25/09] Annual Review of Genetics 29:305-327. IBAMA, 2011. Proposta de Plano Nacional de Fratini, S., L. Ragionieri, C. Papetti, G. Gestão para o uso sustentável do Ca- Pitruzzella, R. Rorandelli, S. Barbaresi, ranguejo-Uçá do Guaiamum e do Siri- and L. Zane. 2006. Isolation and charac- Azul. in: Instituto Brasileiro do Meio terization of microsatellites in Ambiente e dos Recursos Naturais Re- marmoratus (; nováveis . José Dias Neto, organizador. Decapoda; Brachyura). Molecular Ecol- Brasília. 156pp. ogy Notes 6:179–181. Magris, R.A., and R.Barreto. 2010. Mapping Gifford, C.A. 1962. Some Observations on the and assessment of protection of man- General Biology of the Land Crab, grove habitats in Brazil. Pan-American Cardisoma guanhumi (Latreille) in Journal of Aquatic Sciences 5:546-556. South Florida. Biological Bulletin 123:207-223. Mahon, B.C., and J.E. Neigel. 2008. Utility of arginine kinase for resolution of phylo- Govender, Y. 2008. Management Plan Ineffec- genetic relationships among brachyuran tive: Socio-economics determine capture genera and families. Molecular rates of land crab, Cardisoma guanhumi, Phylogenetics and Evolution 48:718– in Puerto Rico. The 93rd ESA Annual 727. Meeting. PS 75.

47

The 14th Symposium on the Natural History of the Bahamas

Miller, M. 2005. Best of the Bahamas. In: Sadovy, Y. and A-M. Eklund. 1999. Synopsis of TravelLady Magazine [online] Available Biological Data on the Nassau Grouper, at: http://www.travellady.com/Issues/ Epinephelus striatus (Bloch, 1792) and June05/1577BestofBahamas.htm the Jewfish, E. itajars (Lichtenstein, [retreived 11/04/11] 1822). In: NOAA technical report NMFS146, A technical report of the Ministério do Meio Ambiente. 2011. Resultado Fishery Bulletin. FAO Fisheries Synop- dos Projetos Habilitados para chamada sis 157. U.S. Department of Commerce, I e II [online] Available at Seattle, WA. http://www.mma. gov.br/sitio/index. php?ido=conteudomonta&idEstrutura=1 Tarazona, J. 1983. A pilot program for the fish- &idConteudo=4707 [retrieved 11/11/11] ery development of shallow areas in the Peruvian upwelling. In: Planning portfo- Nickrent, D. L., W. H. Eshbaugh, and T. K. lio for potential participants, The stock Wilson. 1988. Vascular Flora of Andros assessment collaborative research sup- Island, Bahama. Kendall/Hunt Publish- port program. The University of Mary- ing Company, Dubuque, IA. 221 pp. land Planning Entity. Chesapeake Bio- logical Laboratory, Solomons, MD. Rankine, L.B., G. Seepersad, and R.H. Singh. pp.185-191. 2004. Chapter 11: Profile of the Fishing Industry, The Bahamas. Caribbean Re- Trosper, R.L. 2002. Northwest coast indigenous gional Fisheries Mechanism (Crfm) in: institutions that supported resilience and The Effects of Liberalization and Trade sustainability. Ecological Economics 41: Related Policies on Fisheries and 329-344. Measures Required for their Sustainable Development in the CARIFORUM / Wedes, S. 2004. Cardisoma guanhumi. CARICOM Region. Secretariat, Princess Diversity Web [online] Availble at: Margaret Drive, Belize City, Belize, www.animaldiversity.ummz.umich.edu/ C.A. 105pp. site/accounts/information/Cardisoma_gu anhumi.html. [retrieved 1/25/09] Rodríguez-Fourquet, C. and A.M. Sabat. 2009. Effect of harvesting, vegetation structure Witherington, D. 2009. Land Crab Cardisoma and composition on the abundance and guanhumi. Loxahatchee River District. demography of the land crab Cardisoma Poster Series No. 4. [online] Available guanhumi in Puerto Rico. Wetlands at: www.loxahatcheeriver.org Ecology Management 17:627–640. [retrieved 1/25/09]

Rolle, N. 2007. Crabbing Business Grows In Zhang, D., G. Ding, H. Zhang, and B. Tang. Andros. in: The Bahama Journal, 15 2009. Isolation and Characterization of June 2007. Available at http://www. 10 microsatellite markers in Helice jonesbahamas.com/news/47/ARTICLE/1 tientsinensis (Brachyura: Varunidae). 2993/2007-06-15.html[retreived Conservation Genetics Resources 1:321- 10/05/09] 323

48